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CLINTON NATHAN WOOLSEY
November 30. ~ 904-fanuary ~ 4, ~ 993
BY RICHARD F. THOMPSON
ONE OF THE GREAT achievements of neuroscience in this
century has been characterization of the organization
of sensory en c! motor representations in the cerebral cor-
tex. Rough facts of cortical localization were known from
the nineteenth century en c! earlier, baser! on studies from
brain-ciamagec! humans en c! animals en c! by application of
electrical stimulation. The most important contemporary
scientist to pioneer the fine-grainec! analysis of sensory en c!
motor representations in the cortex was Clinton N. Woolsey.
In the course of his remarkable career he macle many im-
portant en c! funciamental discoveries.
Clinton N. Woolsey was born on November 30, 1904, in
Brooklyn, New York, the son of Joseph Wooc~hull en c! Mathilcia
Louise Aicho~z Woolsey. He left Brooklyn at the age of nine
months ("before cleveloping a Brooklyn accent". He spent
his youth in Orange County, New York, en c! attenclec! a one-
room country school from gracles ~ through 6. He clescribec!
it as an interesting experience, because he "conic! listen to
the lessons given to all the pupils." He attended grades 7
through ~ O in Montgomery, New York, ant! mover! to
Schenevus, New York, for his thirc! year of high school.
There he was awarclec! the H. Bernarc! GoIc! Mecial as the
best student of the year.
361
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362
BIOGRAPHICAL MEMOIRS
In the audience cluring his aware! ceremony was Dr. Walter
AlIan Cowell of Olean, New York, who suggestec! that young
Woolsey move to Olean for his senior year, which he clicI.
Cowell was a physician interested both in the practice of
medicine en c! in research en c! was studying the effects on
diabetes of insulin, which hac! just been cliscoverecI. As a
result of his association with Cowell, Woolsey became greatly
interested in medicine en c! in research. He gracluatec! near
the top of his class en c! spent another year at Olean High
School taking extra work in Latin, French, en c! other subjects.
In 1924 Woolsey enterer! Union College in Schenectady,
New York, where Cowell hac! gracluatecI. Woolsey continues!
his study of Latin en c! French en c! took two years of Greek
en c! the courses neeclec! to enter meclical school. Among
his most impressionable experiences at Union College was
a remarkable psychology professor, Johnny March, who "cle-
scribed the experiments of PavIov and Sherrington so viv-
idly that one felt in the presence of these investigators and
their experimental animals." As a result of this, Woolsey
consiclerec! going to Columbia University for training in
psychology. Instead, he decided to go to medical school,
and was accepted by the Johns Hopkins University School
of Medicine in ~ 928.
During his first year at Hopkins, Woolsey took courses in
histology en c! neuroanatomy from Dr. Marion Hines. At that
time Woolsey intenclec! to become a brain surgeon, so Hines
sent him to work with Dr. Sarah Tower, who was an accom-
plished animal surgeon. Following a special course on To-
calization of function taught jointly by Hines en c! Tower,
Hines invites! Woolsey to work with her on the clog brain,
which led to his first publication: "On the Postural Rela-
tions of the Frontal and Motor Cortex of the Dog" (1933~.
Before finishing his fourth year of medical studies, Woolsey
clevelopec! pulmonary tuberculosis (a not uncommon con
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CLINTON NATHAN WOOLSEY
363
dition for medical students of the day) and had to leave
school for six months to recuperate in a sanitarium. He was
advised that his goal of an internship in surgery was too
physically demanding and might reactivate his pulmonary
lesion. Dr. Philip Bard had just been recruited to Hopkins
from Harvard, and he invited Woolsey to work in his lab.
Woolsey soon realized that his future lay in physiology and
not brain surgery. (In later years, those of us who worked
with him through all-night experiments and 14-hour brain
surgeries on monkeys were amazed at his enormous energy
and robust health. )
During his time at Hopkins, Woolsey married Harriet E.
Runion (in ~ 942) . They had three children: Thomas Allen
Woolsey, M.D., now a leading neuroscientist, John David
Woolsey, M.F.A., an artist and medical illustrator, and Ed-
ward Alexander Woolsey, Ph.D., a zoologist and teacher.
. . .
Woolsey remained at Hopkins in physiology until 194S, when
he accepted his appointment as Charles Sumner Slichter
professor of neurophysiology at the University of Wisconsin
medical and graduate schools in Madison. He remained at
Wisconsin for the rest of his career and life.
I had the great good fortune to work in Clinton Woolsey's
laboratory for four years from 1955 through 1959. I com-
pleted by Ph.D. thesis in 1955-56 in his laboratory (my ma-
jor professor, W. l. Brogden, in the psychology department
did not have facilities for my work, and Woolsey kindly al-
lowed us to use his laboratory). I then spent three years in
Woolsey's laboratory as an NIH postdoctoral fellow. It was a
most exciting environment. Much of the work in the labo-
ratory at the time focused on the organization of the motor
cortex in a series of primates (including chimpanzees) us-
ing electrical stimulation and on the organization of sen-
sory (and polysensory) cortical areas using surface-evoked
potentials. P. W. Davies visited the lab during that time and
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BIOGRAPHICAL MEMOIRS
clescribec! the new extracellular microelectrocle technique
he en c! {erzy Rose hac! clevelopecI. {erzy Rose visitor! one
summer, en c! using this technique we completec! the first
single-unit recording study of the tonotopic organization of
the auditory cortex (in cat) (1960~.
During my time (and, of course, earlier and later, as well)
there were extraorclinariTy taTentec! scientists in the labora-
tory. Konrac! Akert proviclec! solic! expertise in neuroanatomy
(e.g., 1961), Joseph Hint! was expert in the auditory system
en c! all matters acoustic (1960), en c! W. I. Welker en c! Rob
ert Benjamin were young scientists at the height of their
productivity (e.g., 1957~. There were many others as well
(e.g., 1957~. Woolsey was a very tolerant laboratory chief. If
the work we clic! was to some degree relevant to cortical
organization en c! functions en c! was carefully clone, we were
free to follow our own interests. Personally, Woolsey was a
gentle man I never saw him lose his temper. He was an
icleal role moclel in that he was totally focuses! on the work
(anc! his family), was objective, en c! never engages! in act
hominem. However, if you took a particular position on
cortical organization, you hac! better be preparer! to clefenc!
it. He had very high standards and expected the same of
everyone. Morale in Woolsey's laboratory was extremely high.
Woolsey was a superb but infrequent lecturer, often teach-
ing by demonstration. At that time textbooks states! that
complete removal of the neocortex in monkeys causer! vir-
tual paralysis. In the meclical student physiology course
Woolsey once clemonstratec! a fully clecorticate rhesus mon-
key, which he hell! on a stick chain while the monkey chaser!
him arounc! the lectern trying to bite him. This fincling
J ~
was, of course, much more than simply a demonstration.
Travis en c! Woolsey (1956) shower! that after bilateral re
moval of all neocortex in stages, monkeys conic! show con
siclerable recovery of motor function en c! become capable
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CLINTON NATHAN WOOLSEY
365
of locomotion if given adequate postoperative physical
therapy. Recovery of function following brain injury was of
creep interest to Woolsey. I assistec! Woolsey in preparation
of two of the clecorticate macaques (we clic! them in two
stages) en c! in their postoperative care. Woolsey hac! clevel-
opec! a methoc! of subpial surgical aspiration of cortex that
macle it possible to remove TocaTizec! regions without ciam-
age to adjacent regions, or of an entire hemisphere of cor-
tex with minimal bleecling. He was a superb experimental
neurosurgeon.
Woolsey became the Charles Sumner Slichter professor
emeritus at the University of Wisconsin in 1975, but he by
no means retiree! from his work. He publisher! a lancimark
paper on localization in somatic sensory en c! motor areas
of the human cerebral cortex in 1979, en c! until shortly
before his cleath he was hare! at work bringing to comple-
tion his extensive ciata on cortical localization in the chim-
panzee. I participates! in these studies, en c! he caller! me, I
believe in 1990, to check on some cletaiTs.
In the course of his long en c! productive career, Clinton
Woolsey receiver! many honors en c! awards, inclucling Phi
Beta Kappa (1928), the Franklin P. Mall Awarc! in anatomy,
Johns Hopkins University ( 1933), National Academy of Sci-
ences membership (1960), the Mecial of Faculty of Mecli-
cine, Free University, Brussels, Belgium ~ ~ 968), charter
membership in the Johns Hopkins Society of Scholars (1968),
an Sc.D. (honoris cause) from Union College (1968), hon-
orary membership in the Academy of Neurosurgery ~ ~ 973),
honorary membership in the American Neurological Asso-
ciation (1975), en c! the Ralph W. Gerarc! Awarc! from the
Society for Neuroscience, with l. F. Rose (1982~. He server!
on numerous NIH committees en c! was cleeply involves! in
international scientific activities.
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BIOGRAPHICAL MEMOIRS
PROFESSIONAL HISTORY
Wacle Marshall hac! joiner! Philip Barcl's department at
Johns Hopkins in the 1930s en c! hac! worker! with the cath-
ocle ray oscilloscope in Ralph Gerarcl's laboratory. He en c!
Albert Grass at Harvarc! built such equipment for Barcl's
laboratory, en c! Marshall, Woolsey, en c! B arc! undertook the
first cletailec! mapping of the somatic sensory area of the
cerebral cortex of the cat en c! monkey, using the new evokoc!
potential technique (1937, 1941, 1942~. This formed the
basis of much of Woolsey's future work. He mapped the
cortical sensory areas (anc! motor areas) in many mamma-
lian species in great cletail. Woolsey hac! a creep en c! abicling
interest in the comparative clevelopment of functional ar-
eas of the neocortex, an interest he conveyor! to his student
en c! colleague W. I. Welker, who has continues! this impor-
tant tradition to the present. These studies were clone with
great care and attention to detail and led to a general for-
mulation in the late 1940s of the receptotopic organization
of cortical receiving areas, the general plan of which is
largely unchallengec! to this clay.
The power of this comparative approach is clear in the
following passage:
Of particular significance in the evolution of these [somatic sensory mo-
tor] fields is the central position of the hand areas of SI and MI. In the
primates the hand achieves a high degree of corticalization in the precen-
tral and the postcentral fields. Because of the central location of the hand
areas, the simple basic pattern of organization seen in the rodent, where
the parts are represented in relation to one another much as they exist in
the actual animal, apparently becomes distorted in evolution as cortical
representation for the hand increases, with the result that in chimpanzee
and in man the sensory and the motor face areas lose continuity with the
centers for occiput and neck, which remain associated with the trunk rep-
resentations. In macaque this separation of face from occiput has taken
place in the postcentral gyrus, but in the precentral field the motor pattern
still hangs together as it does in lower forms. Evidence for a transitional
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CLINTON NATHAN WOOLSEY
367
status in the postcentral area in the smooth-brained marmoset has been
reported and illustrated elsewhere. That this separation of cortical centers
for face and occiput is not the result of an en bloc reversal of the projec-
tions of the cervical segments upon the cortex as was once suggested (1942),
but rather is due to expansion of the hand area and disruption thereby of
the cortical pattern, is supported by the finding that the trigeminal nerve
projects not only to the lower classical face area but also to the "upper"
head area, where not only the occiput but other parts of the head and face
are represented ( 1958, pp. 65-66~ .
In his initial studies Woolsey focuses! on the somatic sen-
sory cortex, but he quickly extenclec! the work to auditory
en c! visual areas. It was known that regions of the cochlea
responclec! selectively to different tone frequencies, but little
was known about the auditory cortex. Woolsey en c! Walz!
(1942) completec! a technical tour cle force by selectively
stimulating TocaTizec! regions of auditory nerve fibers in the
cochlea en c! mapping the patterns of evokoc! responses on
the auditory cortex of the cat en c! monkey. This was the
first clear demonstration of tonotopic (actually cochicotopic)
organization of the auditory cortex. They follower! this by
examining effects of cochIear lesions on click-evokoc! re-
sponses in the auditory cortex (1946~.
Early in the 1940s Woolsey cliscoverec! the existence of a
seconc! somatic sensory receiving area in the cortex of the
cat, clog, en c! monkey ~ ~ 943) en c! subsequently cliscoverec!
secondary auditory en c! visual areas. Both E. D. Adrian en c!
Woolsey are creclitec! with inclepenclent discovery of the ex-
istence of this seconc! somatic sensory area. Actually, it ap-
pears that Woolsey was first. The following is a quote from
a letter written to me by Clinton's son Thomas:
I believe Dad and his colleagues in Baltimore discovered a second somatic
area independently and about the same time as Adrian. My father rarely
expressed disappointments in others. However, I think this is one case
where he was both very disappointed and surprised. Evidently, early during
the Second World War, Adrian was in the United States and visited the
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BIOGRAPHICAL MEMOIRS
laboratory in Baltimore, Maryland. As Father described it, he [Clinton Woolsey]
spent a long time very carefully explaining his discovery of a second so-
matic area. Father said that Adrian nodded and made comments regarding
the data that he was being shown, but said nothing about his own work,
which Father was greatly surprised to see published several months later. I
think Dad felt betrayed in his confidence. In any case, a review of the data
suggests that Dad provided the first convincing evidence of an orderly
sequence in a second full representation. Adrian's note had only a few
points that were outside the region of what was already known to be the
somatic area (SI)" (Thomas Woolsey, personal communication, Nov. 22,
1994~.
Woolsey en c! associates carefully mapper! the primary vi-
sual area of the cortex, clemonstratec! the cletailec! retinotopic
organization, en c! mapper! a seconc! visual area ~ ~ 946, ~ 950) .
In yet another series of pioneering studies, Woolsey en c!
associates mapper! the somatic sensory projections to the
cerebelIar cortex (1945) en c! the organization of projec-
tions from the cerebral cortex to the cerebellar cortex (1952~.
In still yet another series of pioneering studies, Woolsey
joiner! forces with {erzy Rose to complete a cletailec! lesion
retrograde degeneration mapping of the projections from
the auditory region of the thalamus (meclial geniculate bocly)
to the auditory cortex in light of the physiological organi-
zation of the auditory cortex Woolsey en c! Walz! hac! cle-
finec! earlier (1949~. Rose en c! Woolsey completec! similar
studies on the projections of the meclioclorsal nucleus to
the orbitofrontal cortex (1947) en c! on the relations be-
tween the anterior thalamic nuclei en c! the limbic cortex
(1948~. As notes! by Clinton's son Thomas in the presenta-
tion statement for the Ralph W. Gerard Award to Woolsey
en c! Rose in 1982, these studies clemonstratec! that (~) there
was a direct correspondence between cortical cytoarchitec-
tonic fielcis en c! functionally clefinec! regions of the cortex
(this structure-function concept was under attack at the time),
(2) each functional en c! cytoarchitectonic region of cortex
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CLINTON NATHAN WOOLSEY
369
receiver! a distinctive input from a specific thalamic nucleus
(the concept of thalamotelencephalic clepenclencies), en c!
(3) these connections either conic! be restrictec! or be clis-
tributec! more wiclely to several functional en c! cytoarchitec-
tonic areas (the concepts of essential en c! sustaining projec-
tions) .
In an ongoing series of exquisitely cletailec! studies, Woolsey
en c! associates mapper! the primary en c! supplementary motor
areas of the cerebral cortex, using electrical stimulation in
a wicle range of primates en c! other mammals (1952, 1957,
1958) en c! comparer! sensory en c! motor maps in both pre-
anc! postcentral cortical areas:
It has now been firmly established that the afferent areas are not strictly
afferent nor are the motor areas entirely motor. The afferent areas (SI and
SII; postcentral and "second" sensory) have well-organized motor outflows
which are still functional months after complete removal of the motor
areas of the frontal lobe, while at the same time it appears that afferent
connections to the frontal motor areas exist independently of the parietal
afferent paths (Figure 1~. Thus, the concept that the rolandic region is
indeed a sensorimotor system, as held by pre-Sheringtonian workers, is
reaffirmed, but with the considerable difference that the region is not an
undifferentiated entity but one compounded of a number of distinguish-
able, individually complete, though interrelated, sensory-motor and motor-
sensory representations. These facts appear to us to have important conse-
quences for studies of the role of the cortex in neurological and behavioral
functions, studies which will require the close cooperation of anatomist,
physiologist, and behaviorist, or the mastery of multiple techniques by single
individuals ( 1958, p. 64~ .
It is perhaps fitting to close this review of Clinton Woolsey's
professional history with an example of his work. Figure ~
is reproclucec! from Woolsey ~ ~ 958) . It shows the cletailec!
maps of a portion of the postcentral gyrus of the Macaca
mulatto, comparing the representation of the belly surface
on the left, obtainer! from evokes! potential maps, to the
representation of movements from the same cortical tissue,
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BIOGRAPHICAL MEMOIRS
~N
t;~t,~v\'!
{
~ tofu
/~' /
:
\ ~.~\
/~p
/
/6 ~
/
/ /6~7~/ J
~/
/~1~
,~,~~1 Uj - ~'
.~1~,~8~)
~'
/
/
POSTCENTRAL ~ - ' - POSTCENTRAL
TACTILE ~MOTOR
FIGURE 1: Comparison of postcentral tactile localization pattern with the postcen-
tral motor localization pattern of Macaca mulatto (1958).
elicitec! by electrical stimulation, on the right. The figu-
rines (a style of data representation Woolsey invented and
perfected) in both cases indicate 2-mm steps along the cor-
text The dark region in each figurine on the left is the
region of skin surface that yielcis the maximum amplitucle
evokes! potential at that cortical locus when the skin is lightly
tapped. The dark regions on the figurines on the right are
the cortical loci where least-intensity electrical stimulation
yielcis the minimal movement shown. Note the exquisite
cletail. In the animal on the right (motor map), the cortical
motor area (precentral gyrus) had been removed bilater-
ally, hence there is a well-organized postcentral motor sys-
tem that can function independently of the frontal motor
paths (see quotation just above). Woolsey also notes that
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CLINTON NATHAN WOOLSEY
371
these two maps are clerivec! from different animals, yet they
show remarkable similarities in their patterns of somato-
topical organization.
It wouic! be very remiss of me to conclucle this biography
of Clinton Woolsey without special mention of his wife
Harriet. She was totally supportive of his work en c! career
en c! acceptec! with profounc! goof! nature his increclible work
scheclule, he wouIc! often work all clay en c! night en c! some-
times longer. Clinton was truly most fortunate. We who
worker! with him remember with great fanciness the eve-
nings at their home. Every summer, when the first sweet
corn was ripe in southern Wisconsin, Clinton en c! Harriet
hell! a corn roast for the laboratory at a local park. Perhaps
it was the influence of Harriet en c! Clinton, but somehow
corn has never taster! quite as goof! since.
~ AC~OWLEDGE MOST gratefully the following documents that pro-
vided information, particularly on the early phases of Clinton Woolsey's
life: the autobiographical document dated April 10,1989, that Clinton
Woolsey wrote to the National Academy of Sciences; the original
nomination (circa 1959) to the National Academy of Sciences; the
Ralph W. Gerard Award presentation statement dated November 1,
1982, that Clinton's son Thomas A. Woolsey wrote for the Society
for Neuroscience; and a letter dated November 22, 1994, that Tho-
mas A. Woolsey wrote to me. Finally, I have innumerable personal
experiences from the time I worked in Clinton Woolsey's laboratory
from 1955 to 1959, the high point of my professional career.
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BIOGRAPHICAL MEMOIRS
SELECTED BIBLIOGRAPHY
1937
With W. H. Marshall and P. Bard. Cortical representation of tactile
sensibility as indicated by cortical potentials. Science 85:388-90.
1941
With W. H. Marshall and P. Bard. Observations on cortical somatic
sensory mechanisms of cat and monkey. 7. Neurophysiol. 4:1-43.
1942
With W. H. Marshall and P. Bard. Representation of cutaneous tac-
tile sensibility in the cerebral cortex of the monkey as indicated
by evoked potentials. Bull. fohns Hopkins Hosp. 70: 399-441.
With E. M. Walzl. Topical projection of nerve fibers from local
regions of the cochlea to the cerebral cortex of the cat. Bull.
fohns Hopkins Hosp. 71:315-44.
1943
"Second" somatic receiving areas in the cerebral cortex of cat, dog,
and monkey. Fed. Proc. 2:55-56.
1945
With J. L. Hampson and C. R. Harrison. Somatotopic localization in
the anterior lobe and lobulus simplex of the cerebellum of the
cat and dog. Fed. Proc. 4:31.
i946
With S. A. Talbot and J. M. Thompson. Visual areas I and II of the
cerebral cortex of the rabbit. Fed. Proc. 5:103.
With E. M. Walzl. Effects of cochlear lesions on click responses in
the auditory cortex of the cat. Bull. fohns Hopkins Hosp. 79:309-
19.
1947
With J. E. Rose. The orbitofrontal cortex and its connections with
the mediodorsal nucleus in rabbit, sheep, and cat. Res. Publ. Assoc.
Nero. Ment. Dis. 27:210-32.
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CLINTON NATHAN WOOLSEY
1948
373
With J. E. Rose. Structure and relations of limbic cortex and ante-
rior thalamic nuclei in rabbit and cat. 7. Comp. Neurol. 89:279-
348.
1949
With J. E. Rose. The relations of thalamic connections, cellular
structure and evocable electrical activity in the auditory region
of the cat. 7. Comp. Neurol. 91:441-46.
1950
With J. M. Thompson and S. A. Talbot. Visual areas I and II of the
cerebral cortex of the rabbit. 7. Neurophysiol. 13: 277-88.
1952
Patterns of localization in sensory and motor areas of the cerebral
cortex. In The Biology of Mental Health and Disease, pp. 192-206.
New York: P. Hoeber.
With P. H. Settlage, D. R. Meyer, W. Sencer, T. P. Hamuy, and A.
M. Travis. Patterns of localization in precentral and "supplemen-
tary" motor areas and their relation to the concept of the premotor
area. Res. Publ. Assoc. Nerv. Ment. Dis. 30:238-64.
With J. L. Hampson and C. R. Harrison. Cerebro-cerebellar projec-
tions and somatotopic localization of motor function in the cer-
ebellum. Res. Publ. Assoc. Nerv. Ment. Dis. 30:299-316.
1956
With A. M. Travis. Motor performance of monkeys after bilateral
partial and total cerebral decortications. Am. f. Phys. Med. 35:273-
310.
1957
With W. S. Coxe, J. F. Hirsch, R. M. Benjamin. W. I. Welker, and R.
F. Thompson. Precentral and supplementary motor areas in Ateles.
Physiologist 1:1 9.
With W. I. Welker, R. M. Benjamin, and R. C. Miles. Motor effects
of cortical stimulation in squirrel monkey. J. Neurophysiol. 20:347-
64.
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374
BIOGRAPHICAL MEMOIRS
1958
With H. F. Harlow. Eds. Biological and Biochemical Bases of Behavior.
Madison: University of Wisconsin Press.
With J. E. Rose. Cortical connections and functional organization
of the thalamic auditory system of the cat. In Biological and Bio-
chemical Bases of Behavior, eds. H. F. Harlow and C. N. Woolsey,
pp. 127-50. Madison: University of Wisconsin Press.
Organization of somatic sensory and motor areas of the cerebral
cortex. In Biological and Biochemical Bases of Behavior, eds. H. F.
Harlow and C. N. Woolsey, pp. 63-81. Madison: University of
Wisconsin Press.
1960
With J. E. Hind, J. E. Rose, P. W. Davies, R. M. Benjamin, W. I.
Welker, and R. F. Thompson. Unit activity in the auditory cortex.
In Neural Mechanisms of the Auditory and Vestibular Systems, eds. G.
L. Rasmussen and W. F. Windle, pp. 201-10. Springfield, Ill.: Charles
C. Thomas.
Organization of the cortical auditory system: A review and a synthe-
sis. In Neural Mechanisms of the Auditory and Vestibular Systems, eds.
G. L. Rasmussen and W. F. Windle, pp. 165-80. Springfield, Ill.:
Charles C. Thomas.
1961
With K. Akert, R. A. Grusen, and D. R. Meyer. Kluver-Bucy syn-
drome in monkeys with neocortical ablations of temporal lobe.
Brain 84:480-98.
1979
With T. C. Erickson and W. E. Gilson. Localization in somatic sen-
sory and motor areas of human cerebral cortex as determined by
direct recording of evoked potentials and electrical stimulation.
7. Neurosurg. 51:476-506.
Representative terms from entire chapter:
auditory cortex